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PubMed2003

Anisotropic representation of forelimb position in the cerebellar cortex and nucleus interpositus of the rat.

The relationship between the spatial location of limb and the activity of cerebellar neurons has received little attention and its nature still remains ambiguous. To address this question we studied the activity of Purkinje and nucleus interpositus cells in relation to the spatial location of rat forelimb. A computer-controlled robot arm displaced the limb passively across 15 positions distributed on a parasagittal plane. The limb was upheld for 8 s in each position, which was identified by the Cartesian coordinates of the forepaw. We selected the neurons whose activities were significantly modulated by forepaw position and found that the majority represented preferentially one spatial dimension of the Cartesian plane both in the cerebellar cortex and nucleus interpositus. In particular, the antero-posterior axis was best represented in cerebellar neuronal discharges. This result suggests that the intermediate part of the cerebellum might encode limb position by way of an anisotropic representation of the spatial coordinates of the limb end-point.

Action Potentials↗
PubMed1997

Cutaneous reflex activity of the cat forelimb during fictive locomotion.

To investigate mechanisms of rhythmic modulation of cutaneous reflex by the central pattern generator for locomotion, we made intracellular recordings from elbow flexor motoneurons, biceps brachii (Bi) and brachialis (Br), and quantitatively analyzed excitatory postsynaptic potentials (EPSPs) evoked by stimulation of superficial radial (SR) nerves during forelimb fictive locomotion, which was evoked in immobilized, decerebrate cats with the low thoracic spinal cord transected. We found that (1) in both Br and Bi, SR stimulation evoked trisynaptic EPSPs (segmental latencies were 2.10 +/- 0.39 ms for Br and 2.07 +/- 0.93 ms for Bi), (2) the SR-EPSPs of Br and Bi were rhythmically modulated with a similar pattern; the maximal and minimal EPSPs appeared in the flexor and extensor phase, respectively, (3) however, there were differences as follows. In the control state where fictive locomotion was absent, the mean amplitude of SR-EPSPs of Br (4.65 +/- 2.76 mV, n = 14) was much larger than that of Bi (1.25 +/- 1.22 mV, n = 10). During fictive locomotion, the maximum amplitude of SR-EPSPs of Br was 5.63 +/- 1.79 mV (n = 18), and the minimum was 3.82 +/- 1.69 mV (n = 18); that is, the maximum during fictive locomotion was larger or smaller than that of the control, while the minimal were always smaller. In contrast, SR-EPSPs of Bi were modulated over the control level; the maximum was 3.97 +/- 1.71 mV (n = 16) and the minimum was 2.24 +/- 1.11 mV (n = 16), both the maximum and the minimum during locomotion being larger than that of the control. These results may suggest that two mechanisms are involved in the rhythmic modulation, first, cyclic facilitation in the flexor phase, and secondly, cyclic inhibition in the extensor phase. The former was involved in both cases of Bi and Br motoneurons, while the latter would be involved only in the case of Br motoneurons.

Animals↗
PubMed1997

Ethanol consumption following recovery from unilateral damage to the forelimb area of the sensorimotor cortex: reinstatement of deficits and prevention of dendritic pruning.

Unilateral injury to the forelimb-representation area of the sensorimotor cortex (FL-SMC) in adult rats results in use-dependent proliferation of dendritic processes, followed by partial pruning, of layer V pyramidal neurons of the contralateral homotopic cortex. In development, 'exuberant' growth of neurons is often followed by pruning, a process that has been associated with a glutamatergic-NMDA receptor mechanism. A related mechanism may play a role in injury-related pruning of dendrites in adults. The N-methyl-D-aspartate (NMDA) receptor antagonist MK801, administered throughout the pruning phase to adult animals with FL-SMC lesions, prevents dendritic pruning and disrupts behavioral recovery. Ethanol (ETOH) also acts as an NMDA receptor antagonist. It has been shown to reduce NMDA-active ion currents, inhibit NMDA-evoked electrophysiological responses, and decrease glutamate-binding in the hippocampus and cortex. ETOH also affects neuromorphology in the developing and adult cerebellum, hippocampus, and cortex. Ethanol's involvement with NMDA receptor function and its influence on dendritic morphology led us to examine its effect on dendritic pruning and behavioral recovery following unilateral FL-SMC lesions. Lesioned animals were exposed to moderate doses of ethanol in a liquid diet only during the period of dendritic pruning. As with MK801, ETOH prevented pruning and reinstated chronic behavioral asymmetries.

Animals↗
PubMed2003

Reconstructed bone end loads on the canine forelimb during gait.

The objective of this work was to determine bone loading conditions that, when applied to a finite element model, would best reproduce the in vivo strain field as measured by surface-mounted strain rosettes. The present study adopts the basic mathematical approach to load reconstruction introduced by Weinans and Blankevoort (J. Biomech. 28 (1995) 739) who determined the relationship between applied loads and bone strain distribution using ex vivo calibration testing. Our method eliminates the need for subsequent ex vivo calibration tests by instead substituting a computational calibration procedure. This first application of the method is with in vivo strains on the canine forelimb during gait (Coleman et al., J. Biomech. 35 (2002) 1677), but with further refinements the method could be used to reconstruct the in vivo loading conditions in living subjects.

Animals↗
PubMed2002

Forelimb muscle activity following nerve graft repair of ventral roots in the rat cervical spinal cord.

Current research on the cellular mechanisms of nerve regeneration suggests the application of nerve growth factors at the repair sites to be beneficial. To test the effectiveness of this approach, we performed transections of the C6 and C7 ventral rootlets from their original sites in the spinal cord of 18 rats. We investigated the electrophysiological changes in three groups of rats operated on by different repair strategies. Six rats comprised the control group (G1). In the other 12 rats, 24 rootlets were implanted into the spinal cord by means of an intercostal nerve graft through the pia mater immediately after transection. Six rats (G2) had fibrin glue applied at the incision. The last 6 rats (G3) had grafts with acidic fibroblast growth factor (aFGF) added to the fibrin glue. The rats' functional recovery was evaluated electrophysiologically at 6 weeks and 6 months after the operation. Needle electromyography showed profound fibrillation potentials (Daube's scoring system) in the deltoid, biceps, and triceps of the operated forelimbs in all groups 6 weeks after the operation. After 6 months, there was a significant decrease in the amount of fibrillation potentials in all groups (G1, G2 and G3, p < 0.0001, 0.0001, 0.0009, respectively, generalized estimating equation, repeated measures) and a significantly high probability for motor units present in sampled muscles of G2 and G3 as compared to G1 (log odds ratio in G2 = 51.8316, G3 = 57.4262, generalized estimating equation). We conclude that several cervical roots can regenerate through intercostal nerve grafts applied using fibrin glue. Adding aFGF may increase the efficacy of sprouting.

Animals↗
PubMed2004

The central pattern generator for forelimb locomotion in the cat.

This chapter addresses the neuronal organization of the central pattern generator (CPG) for forelimb locomotion in the cat. The focus is on fictive locomotion, as induced by repetitive stimulation of the cervical lateral funiculus (CLF) in the decerebrate preparation. The stimulus time-locked responses of cervical motoneurons (MNs) that follow each current pulse of such repetitive stimulation include disynaptic EPSPs, trisynaptic EPSPs and IPSPs, and polysynaptic PSPs. Their amplitudes are all phase-related to selected aspects of the locomotor rhythm, as determined by the pattern of discharge in nerves to selected muscles. Correlation analysis reveals rhythmic modulation between the responses of extensor and flexor MNs, thereby suggesting mutual interactions between pathways mediating the CLF effects on MNs. It is argued that the shortest CLF pathway to MNs via the CPG is disynaptic: i.e., with only a single intercalated interneuron (IN) between the descending command axons and the MNs. A second proposal is that such intercalated INs contribute to interacting, reverberating IN circuits (i.e., the CPG) through mutual excitation.

Animals↗
PubMed1989

Distribution of recurrent inhibition in the cat forelimb.

This chapter reviews experiments on the distribution of recurrent pathways from motor axon collaterals to alpha motoneurones in the brachial enlargement of the cat. In anaesthetized cats intracellular recording from identified forelimb motoneurones was used to reveal the pattern of recurrent inhibition or excitation following stimulation of muscle nerves. The recurrent connections of the motor nuclei acting on the elbow follow the tight mechanical agonism of the muscles involved. Extensive bidirectional recurrent inhibitory connections were found between motor nuclei innervating elbow and wrist muscles. It is suggested that one group of these connections supports the organization of limb extension, the other group organization of limb flexion. The supinator and the pronator teres motornuclei have identical recurrent connections. Co-convergence from elbow flexor and extensor motornuclei in one and the same motoneurone is frequent. It is suggested that this pattern may serve the stabilization of the radio-ulnar plane. Neither homonymous nor heteronymous recurrent actions were observed in the radial motornuclei acting on the digits, which suggests a lack of recurrent axon collaterals in these nuclei. These results draw attention to the fact that recurrent inhibition is not evenly distributed between all limb motor nuclei.

Animals↗
PubMed1989

Control of forelimb muscle activity by populations of corticomotoneuronal and rubromotoneuronal cells.

We review and synthesize evidence on the activity of corticomotoneuronal (CM) and rubromotoneuronal (RM) cells and single motor units in forearm muscles in monkeys performing alternating wrist movements. The CM and RM cells were identified by post-spike facilitation of rectified forelimb EMG activity. RM cells facilitated more muscles per cell (mean: 3.0 of 6 synergist muscles) than CM cells (2.4/6). Both groups had "reciprocal" cells which also suppressed antagonists of their facilitated target muscles. Unlike CM cells, some RM cells cofacilitated flexor and extensor muscles (5.8 or 12 muscles). During performance of a standard ramp-and-hold force tracking task the firing patterns of CM and RM cells, as well as single motor units, fell into distinct response types. Each population had phasic-tonic and tonic cells. Unique to the CM population were cells whose discharge increased during the static hold period; unique to RM cells were bidirectionally responsive and unmodulated neurons. Many motor units showed decrementing discharge. To estimate the ensemble activities of these populations the response histograms of different cells were summed (with force ramps aligned) in proportion to the relative frequency of each cell type. The population response histogram of CM cells was phasic-tonic, consistent with the predominant response type. The population response of RM cells was also phasic-tonic, but showed a shallower phasic modulation relative to discharge that was sustained during both directions of movement. The population histogram of motor units of a muscle was proportional to the average of rectified multiunit EMG, and typically exhibited decrementing activity during the static hold. The effects of excitatory postsynaptic potentials (EPSPs) on firing probability of motoneurons previously documented in intracellular studies are combined with the mean firing rates in the population histograms and the known amplitudes of CM-EPSPs and RM-EPSPs to infer the relative contributions of the supraspinal cells to tonic discharge of active motoneurons. This analysis suggests that for intermediate levels of force, the CM cells would increment motoneuron discharge by about 9 impulses/second (i.p.s.) and RM cells by about 2.4 i.p.s. The analysis also reveals differences in the population activity of CM and RM cells compared to their target motoneurons, which may be due to other input cells and to recruitment properties of motoneurons.

Animals↗
PubMed1997

The control of forelimb movements by intermediate cerebellum.

In a series of studies, the functional organization of cerebellar regions contributing to the control of forelimb movements via the rubro- and corticospinal tracts has been characterized in the cat. The system consists of the cerebellar cortical C1, C3 and Y zones and their efferent intracerebellar nucleus, the interpositus anterior. Based on analyses of cutaneous and muscle afferent climbing fibre input, of corticonuclear connections and of limb movements controlled, a modular organization of this cerebellar control system is proposed. Each module consists of a number of cortical microzones, defined by their homogeneous climbing fibre input, and a group of neurones in nucleus interpositus anterior on which these microzones converge. The input to climbing fibres is multi-modal and originates from cutaneous A beta (tactile), A delta and C (nociceptive) fibres and from muscle afferents. The cutaneous receptive fields have spatial characteristics suggestive of a relation to elemental movements. For most climbing fibres, the spatial relationship between cutaneous and muscle afferent input is such that the muscle afferent input originates from muscles that, if activated, would tend to move the cutaneous receptive field of the climbing fibre towards a stimulus applied to the skin. By contrast, the limb movement controlled by the module often has the opposite direction, and would thus tend to move the cutaneous receptive field away from a stimulus applied to the skin. Functional implications of this organization for the involvement of these regions in acute and adaptive motor control of limb movements are discussed.

Afferent Pathways↗
PubMed1985

Effects of unilateral and bilateral training in a reaching task on dendritic branching of neurons in the rat motor-sensory forelimb cortex.

Effects of motor training on a neocortical nerve cell population involved in performance of the motor task were assessed by measuring Layer V pyramidal neuron apical dendritic branching in motor-sensory forelimb cortex of rats trained to reach into a tube for food. Rats were trained to reach with the forepaw they preferred to use (group PRAC), the nonpreferred forepaw (REV), both forepaws (ALT), or neither forepaw (CONT). Across groups, hemispheres opposite trained forepaws had larger apical dendritic fields, in terms of total dendritic length, number of oblique branches from the apical shaft, and length of terminal branches. Similar, although somewhat less consistent, effects were seen when results were analyzed for between- (CONT vs ALT) and within-subject comparisons (trained vs nontrained hemispheres of REV and PRAC). This finding is compatible with the hypothesis that altered dendritic patterns, with associated synapses, are involved in storage of information from the training experience. The within-subject effects mitigate suggestions that these differences arise from generally acting hormonal or metabolic consequences of the training experience, although the possibility that these effects result from neural activity per se and are unrelated to information storage cannot be excluded.

Animals↗
PubMed1997

Muscle activity during forelimb reaching movements in rats.

The absolute timing, and correlation in time of activity in a number of shoulder, elbow and wrist muscles in the rat was analysed in relation to onset and termination of the extension phase of a skilled forelimb reaching movement. Movement onsets were analysed separately in the component upward, forward and medial directions. On average, movements in the upward and medial directions occurred together, prior to forward movement. Latissimus dorsi activity was the earliest muscle event, occurring approximately 150 ms prior to movement, whilst onset of teres major activity, possibly related to paw elevation, had the highest temporal correlation with movement onset. Triceps activity was strongly time locked to the end of the reach, and may have provided final extensor thrust to complete the reaching movement. Although it is possible to speculate on possible roles of particular EMG bursts, multiple (often anatomically antagonistic) muscles were active at all stages of the reach, and no unique muscle relationships to initiation of individual movement components could be identified. It is concluded that reaching movements are produced by temporal variation in distributed activity among all available muscles.

Animals↗
PubMed1997

Corticosubthalamic input zones from forelimb representations of the dorsal and ventral divisions of the premotor cortex in the macaque monkey: comparison with the input zones from the primary motor cortex and the supplementary motor area.

Employing double anterograde axonal tracing in combination with intracortical microstimulation, we examined the distribution patterns of corticosubthalamic inputs from forelimb representations of the dorsal (PMd) and ventral (PMv) divisions of the premotor cortex in the macaque monkey. The inputs from the PMd and PMv were distributed mainly in the medial aspect of the subthalamic nucleus (STN), in which their distribution areas overlapped each other. By the same experimental approach, we further compared corticosubthalamic input zones from the PMd/PMv with those from the primary motor cortex (MI) and the supplementary motor area (SMA). The input zones from the PMd/PMv and SMA largely overlapped in the medial aspect of the STN, whereas the input zones from the PMd/PMv and MI were virtually segregated mediolaterally in the STN.

Animals↗
PubMed1998

At low doses, harmaline increases forelimb tremor in the rat.

A behavioral preparation especially sensitive to low-dose drug effects on fine motor behavior in rats was used to assess the tremorogenic effects of harmaline, an indole alkaloid and beta-carboline derivative. Rats that were trained to press downward on a force transducer for water reinforcement were initially administered harmaline (0.5 and 1.0 mg/kg) in an acute dosing regime. Immediately following the day of initial acute exposure to 1 mg/kg, 3 consecutive days at this dose ensued, providing for a 4-day, repeated-dosing analysis. Harmaline did not significantly suppress task engagement during either acute or repeated dosing. Acute administration of harmaline dose-dependently increased power in the high-frequency (10-25 Hz) band of the power spectrum (tremor) without affecting overall forelimb force output. Upon continued administration, tremor remained significantly elevated above vehicle values. Harmaline also slowed the rats' licking frequency, an effect that did not diminish with repeated dosing. Harmaline increased the durations of individual responses during acute dosing and continued to exert this effect with repeated dosing. The effects reported in the present study may represent low-dose harmaline-induced alterations in the olivo-cerebellar system.

Animals↗
PubMed1999

Neuronal activity in rat red nucleus during forelimb reach-to-grasp movements.

The activity of 259 task-related red nucleus neurons was recorded using chronic electrophysiological methods in free moving rats. Modulations in activity were analysed in relation to onset (first detected wrist movement) and end (arrival of the paw over the food) of a reach-to-grasp movement. Excitatory peaks were found to begin before, during and after the reach, but there were clear peaks in the distribution of onset times after reach-onset and before reach-end, reflecting the fact that one third of all peaks began specifically during the reach, although this occupied only a small fraction of the analysis time. Both excitations and inhibitions showed a strong tendency to end in close temporal association with reach-end. Analysis of excitatory modulation amplitudes showed that the largest peaks were formed when data was aligned to reach-end, and that these largest peaks nearly all began during the reach and ended precisely at the time the paw would have been about to grasp the food. The spread of neural activation onset times throughout the course of the complex reach-to-grasp movement is consistent with a relationship of individual neurons in the rat red nucleus with movements of all parts of the forelimb, as would be expected if all limb muscle groups are represented in the nucleus. On the other hand the disproportionate number of modulations that occur during the reach and their strong alignment with time of reach-end suggests there is a bias in red nucleus function towards the control of distal motions associated with accurate grasp, consistent with the result of recent lesion studies. This provides indirect evidence that functionally the rat red nucleus may be organized in a similar way to that of monkeys, in which an important role in control of accurate distal movements is well established. The possibility is discussed that red nucleus offers a timing signal for co-ordination of movements across joints, in particular the precise distal-proximal binding normally seen in accurate reach-to-grasp movements.

Action Potentials↗
PubMed1981

Mapping of forelimb afferents to the cuneate nuclei of the rat.

Stumps of rat forelimb peripheral nerves were soaked in horseradish peroxidase solution. Impregnation of the median, ulnar, radial, and musculocutaneous nerves revealed that each nerve had definite projection zones in the cuneate nuclei. The radial nerve projected quite heavily to the lateral cuneate nucleus and to a narrow band along the dorsal edge of the main cuneate nucleus. Staining of the deep radial and superficial radial nerves showed that they projected heavily to the lateral and main cuneate respectively. The ulnar and median nerves projected primarily to medial and lateral portions of the main cuneate nucleus, ventral to the terminal field of the radial nerve. The rostral and caudal portions of the main cuneate nucleus were discriminable by cytoarchitectural characteristics and by terminal fibre orientation.

Afferent Pathways↗
PubMed1996

The placement of a striatal ibotenic acid lesion affects skilled forelimb use and the direction of drug-induced rotation.

The motor consequences of excitotoxic striatal damage have been evaluated extensively in the rat, using tests of whole body motor asymmetry and of deficits in skilled paw and limb movements. However conflicting results of both the type and extent of behavioural deficits have been reported, particularly in the direction of rotation observed in response to the dopamine receptor agonist, apomorphine. The present study investigated the effect of unilateral ibotenic acid lesions in the dorsal striatum of the adult rat, placed at either anterior, posterior, medial, or lateral loci, on rotation in response to both amphetamine and apomorphine, and in the "staircase test" of skilled forelimb use. In a 2 x 2 matrix design experiment, adult female albino rats received a double unilateral lesion of 0.5 microliter 0.06 M ibotenic acid injected at each of two sites either anterior (medial and lateral), posterior (medial and lateral), medial (anterior and posterior), or lateral (anterior and posterior). Rats that received posterior lesions showed a marked ipsilateral rotation in response to both amphetamine and apomorphine, while animals receiving anterior lesions showed little ipsilateral or a slight contralateral bias. Rats receiving lateral lesions showed a marked impairment of contralateral paw use on the "staircase test," while animals with medial lesions showed no significant difference to control unoperated animals. These results confirm the somatotopic organisation of the dorsal striatum in its control of motor functions, and indicate the need to take into account the locus of an excitotoxic lesion in the design of lesion and transplantation studies if we are to achieve reliable tests of the behavioural deficits and recovery.

Animals↗
PubMed1996

Functional structuration of the cerebellar efferent network involved in forelimb control, during processing of an associative conditioning.

A simplified sensorimotor pathway was subjected to an associative conditioning in fully awake cats. It was defined by the location of electrodes for applying the conditional stimulation (CS) in the cerebellar interpositus nucleus (IN) and the unconditional stimulation (UCS) on the skin of the forelimb. Electrodes chronically implanted all along the pathway allowed the simultaneous study of the motor and the neurobiological effects of conditioning. Persistent motor changes were observed in parallel with interposito-rubral and thalamocortical synaptic efficiency changes. The two kinds of event evolved similarly and significant correlations were found between them. Changes appeared to be induced according to modalities similar to those underlying elementary mechanisms of neuronal plasticity. The modulations of thalamocortical synaptic efficiency inside the primary motor cortex (MI) determined a new pattern of distribution of cerebellar effects on musculature, indicating that in adult brains the functional organization of central motor pathways is not definitely rigid it can still be subject to adaptive modifications.

Animals↗
PubMed1978

Habituation of the forelimb-withdrawal response in neonatal rats.

The purpose of the present research was to examine the ontogeny of habituation in the neonatal rat, using the forelimb-withdrawal response. Thresholds and latencies of the response, changing patterns of responding to shock stimuli, and habituation of the response were studied in rats 3 to 15 days of age. It was found that although response thresholds do not change during this period of development, response latencies decrease and amplitudes increase. Three- and 6-day old pups remain active much longer following shock than do 10- and 15day-olds. When compared for habituation to different frequencies of stimulation, 3-day-old rat pups were found to be much more susceptible to habituation at low frequencies than 15-day-olds. Insertion of a simple intense shock in the habituation series produced marked dishabituation in the 3-day-olds but little or none in the 15-day-olds. This pattern of results was obtained regardless of the locus of the dishabituating shock. Analysis of response latencies showed that although 15-day-olds responded 2 to 3 times more quickly than 3-day-olds, both age groups responded more rapidly to a strong shock than to a weak one. During habituation, response amplitudes of both 3- and 15-day-old pups declined with no change in latency. Age-related differences in habituation were shown to be independent of differences in reactivity to shock and more likely due to the emergence of response sensitization.

Age Factors↗